Lasallegrion koebelei Crawford, 1912

Janšta, Petr, Delvare, Gérard, Baur, Hannes, Wipfler, Benjamin & Peters, Ralph S., 2020, Data-rich description of a new genus of praying mantid egg parasitoids, Lasallegrion gen. n. (Hymenoptera: Torymidae: Podagrionini), with a re-examination of Podagrion species of Australia and New Caledonia, Journal of Natural History 54 (9), pp. 755-790 : 773-779

Lasallegrion koebelei Crawford, 1912 , comb. n.

( Figures 5b , 6 a–e, 7 a, 8 , 9 , 10 a–c, 13 a–b)

Podagrion koebelei Crawford, 1912: 4–5 ,

Podagrion grotii Girault, 1915: 291–292 .

Podagrion dolichurum Cockerell, 1930: 2–3 , Podagrion holbeini Girault, 1923: 8 ;

Podagrion metatarsum Girault, 1929: 341–342 ;

Figure 2 ; ♀ holotype (examined), AustraliaSouth Australia ( USNM). Label: 606, Australia Koebele, Type No. 14342 U.S.N.M., Podagrion koebelei ♀ Type, Cwfd. Bouček (1988): 141.

♀ holotype (examined), Australia – Queensland ( QMB); Label: HOLOTYPE Hy. 3319 , E.C.D. 1983, Photographed Specimen, Podagrion grotii Girault ♀ Type, ENTI 6.12. Bouček (1988): 141 (synonymy under P. koebelei ).

Figures 1–5 . ♀ holotype (examined), New Caledonia ( AMNH); Label: Podagrion dolichurum, Ckll . TYPE, Noumea , New Caledonia. Bouček (1988): 141 (synonymy under P. koebelei ).

syntypes (examined), Australia – Victoria ( QMB); Label: SYNTYPES T. 5090 E.C.D. 1983 , Podagrion holbeini Gi. ♀ ♂ Types, Brigth Bred from egg capsule of large mantis, Photographed specimen. Bouček (1988): 141. Syn. n.

syntypes (examined), Australia – Victoria ( QMB); Label ( QMB): SYNTYPES T. 5097 E.C.D. 1984 , Photographed specimen, Podagrion metatarsum ♀ ♂ Gi. Type. Bouček (1988): 141. Syn. n.

Non–type material examined

Australia, ATC, Canberra , National Botanical Garden, 35.279831°S 149.110501°E, 575 m a.s. l., 5. Feb. 2014, em. Feb.–Mar. 2014, ex Archimantis sp. ootheca, leg. P. Janšta (7 ♀♀, lc_0103, lc_05–08, ZFMK; 6 ♀♀, lc_33–34, lc_43–44, lc_46, lc _ 55, CUPC; 4 ♀♀, GDEL3175 GDEL3177 , lc_04, CBGP); Australia, ATC, Canberra, Black Mtn. , 1.–14. Dec. 2001, leg. NA (1 ♀, lc _ 09, ANIC); Australia, ATC, Canberra , Black Mtn., 35.16°S, 149.06°E, 1.–14. Feb. 1999, Malaise trap, leg. G. Gibson (1 ♀, lc _ 10, ANIC); Australia, ATC, Canberra , Black Mtn., 35.16° S, 149.06°E, 15.–28. Feb. 1999, Malaise trap, leg. G. Gibson (1 ♀ lc _ 45, ANIC); Australia, ATC, Canberra , Black Mtn., Dec. 1982, Malaise trap, leg. I.D. Naumann & J.C. Cardale (1 ♀, lc _ 36, ANIC); Australia, ATC, Canberra, Apr. 1948, ex mantid ootheca, ex ethanol, leg. NA, (1 ♀, lc _ 37, ANIC); Australia, ATC, Tharwa , coll. 28. Feb. 1988 emg. 1. Mar. 1988, ex ootheca Archimantis sp., leg. J. Balderson, (1 ♀♀, lc_40, ANIC); Australia, QLD, Hann River , 15.11°S 143.52°E, 18. Dec. 1993– 14. Jan. 1994, Malaise trap, leg. P. Zborowski & Edwards, (1 ♀, lc _ 29, ANIC); Australia, QLD, Hann River , 15.11°S 143.52°E, 20. Mar.–24. Apr. 1994, Malaise trap, leg. P. Zborowski & G. Turner, (1 ♀, lc _ 31, ANIC); Australia, QLD, 13 km E. by S. of Weipa, 12.40 S, 143.00E, 16. Jan.–16. Feb. 1994, Malaise trap, leg. P. Zborowski & D. Khalu, (1 ♀, lc _ 30, ANIC); Australia, QLD, 13 km E. by S. of Weipa, 12.40 S, 143.00E, 15. Nov.–16. Dec 1993, Malaise trap, leg. P. Zborowski, (1 ♀, lc _ 32, ANIC); Australia, NSW, Whiskers , 7 km W NW of Hoskinstown, 35.24 S, 149.23E, 8. Jan. 1993, leg. M. S. Upton, (1 ♀, lc _ 35, ANIC); Australia, NSW, Armidale , Sep. 1991, ex mantid ootheca, leg. H. Coombs, (1 ♀, lc _ 38, ANIC); Australia, NSW, 4.1 km W of Williamsdale , 19. Dec. 1994, on Eucalyptus flower, leg. G. Maynard & G. Davis, (1 ♀, lc _ 41, ANIC); Australia, ATC, Canberra, National Botanical Garden , 35.279831°S 149.110501°E, 575 m a.s.l., Feb.–Mar. 2014, ex Archimantis sp. ootheca, leg. P. Janšta (4 ♂♂, lc_11–14, CUPC; 3 ♂♂, lc_50–52, ZFMK); Australia, SE Queensland: Tambourine Mts ., 11.–18.iv.1935, R.E.Turner, B.M. 1935–240, ♀ Podagrion koebelei Crawf. , det. Z. Bouček 1978 (1 ♀, lc _ 74, ANIC); Australia: QLD, 0.5 km S Gordonvale , 10 m, 27.iv.1990, J. Heraty, H90/036, edged sugarcane field, Univ. Calif. Riverside Ent. Res. Mus. UCRC ENT 78401 (1 ♀, lc _ 75, UCRC); Australia: QLD, Munduberra , 13.iv.–12.v.2000, C. Freebairn, MT, Univ. Calif. Riverside Ent. Res. Mus. UCRC ENT 147599 (1 ♀, lc _ 76, UCRC); Australia: SA, Belair NP, gate 9 MT, 21.–27.i.2008, J.T. Jennings (1 ♀, lc _ 77, SMNS); Australia: SA, Kangaroo Island, Flinders Chase NP, Gosselands , 35.93325´S, 136.9326´E, 16.i.2019, Janšta, Böhmová, ex Archimantis sp. ootheca, em. ii.–iv. 2019, PJ19020_8_01, in EtOH (20 ♀♀, lc_78–lc _ 97, CUPC); Australia: SA, Kangaroo Island, Flinders Chase NP, Gosselands , 35.93325´S, 136.9326´E, 16.i.2019, Janšta, Böhmová, ex Archimantis sp. ootheca, em. ii.–iv. 2019, PJ19020_8_2 (10 ♀♀, lc_98107, CUPC; 10 ♀♀, lc_108–117, CBGP); Australia: SA, Kangaroo Island, Flinders Chase NP, Gosselands , 35.93325´S, 136.9326´E, 16.i.2019, Janšta, Böhmová, ex Archimantis sp. ootheca, em. ii.–iv. 2019, PJ19020_8_3 (6 ♀♀, lc_118–123, CUPC; 6 ♀♀, lc_124–129, ZFMK); Australia: SA, Kangaroo Island, Flinders Chase NP, Gosselands , 35.93325´S, 136.9326´E, 16.i.2019, Janšta, Böhmová, ex Archimantis sp. ootheca, em. ii.–iv. 2019, PJ19020_8_4, homeomorph males (3 ♂♂, lc_130–132, CUCP); Australia: SA, Kangaroo Island, Flinders Chase NP, Gosselands , 35.93325´S, 136.9326´E, 16.i.2019, Janšta, Böhmová, ex Archimantis sp. ootheca, em. ii.–iv. 2019, PJ19020_8_5, heteromorph males (4 ♂♂, lc_133–136, CUCP; 5 ♂♂, lc_137–141, CBGP; 5 ♂♂, lc_142–146, ZFMK); Australia: SA, Kangaroo Island, Flinders Chase NP, Gosselands , 35.93325´S, 136.9326´E, 16.i.2019, Janšta, Böhmová, ex Archimantis sp. ootheca, em. ii.–iv. 2019, PJ19020_8_7, in EtOH heteromorph males (10 ♂♂, lc_147–156, CUCP); Australia: SA, Kangaroo Island, Flinders Chase NP, Gosselands , 35.93325´S, 136.9326´E, 16.i.2019, Janšta, Böhmová, ex Archimantis sp. ootheca, em. ii.–iv. 2019, PJ19020_8_8, in EtOH (3 ♀♀, lc_157, lc _ 159, 2 ♂♂, lc_160161, ZFMK), card-mounted (1 ♀, lc _ 158, ZFMK).

Diagnosis. Antenna ( Figure 6b ) with all funicular segments slightly or distinctly longer than broad, length of segments decreasing from proximal to distal, F7 at least 1.1× as long as wide. Combined length of pedicel and flagellum 1.45–1.65× breadth of head. Antenna inserted high on the head, distance from lower edge of toruli to ventral margin of clypeus 1.1–1.6× as long as distance from lower edge of toruli to anterior ocellus. Parascrobal area remarkably raised above the outline of head anteriorly (best seen in dorsal or lateral view) and scrobal depression deep with interantennal process long, raised slightly above outline of parascrobal area ( Figure 8a ). Head height 0.85–1.02× as long as marginal vein. Notauli almost parallel posteriorly ( Figure 8d – indicated by arrows). Propodeum with adpetiolar area (behind posterior branches of carinae) coarsely rugose. Metafemur large, 1.88–2.73x as long as broad, with five long ventral teeth plus one composite terminal one, at least length of 3 rd and 4 th tooth same as or longer than width of tibia opposite to tooth ( Figure 8e ). Costal cell of fore wing with two complete rows of setae on the underside ( Figure 8f ). Ovipositor very long, between 1.6 and 2.5x as long as body ( Figure 6a ).

Additional characters. FEMALE (N = 90): Body length excluding ovipositor between 3.3 mm and 4.4 mm; length of ovipositor 5.3–9.8 mm.

Colour. Head, meso- and metasoma entirely metallic bronze blue to green with coppery reflections. Scape, pedicel and flagellum dark brown. Pro-, meso- and metacoxa and outer side of metafemur metallic bronze blue to green with coppery reflections, inner median areas of pro- and metacoxa brown, paramedian parts yellow. Median areas of outer side of pro- and mesofemur and of inner side of metafemur brown with coppery reflections, paramedially yellow. Pro-, meso- and metatibia yellow. All tarsi yellow but pretarsi brown. Ovipositor pale yellow to white, sheaths brown. Fore wing hyaline, wing venation pale to slightly brown, setae brown.

Head. Head 1.15–1.28x as broad as high ( Figure 8b ) and 1.6–2.2x as broad as long ( Figure 8a ); 1.17–1.32x as broad as pronotum breadth. Frontovertex 0.83–0.96x as broad as eye height. Eye 1.16–1.35x as high as long. Parascrobal protuberance 0.04–0.11x as high as head height. Malar space 0.44–0.6x as long as breadth of oral fossa and 0.28–0.37x as long as eye height. Clypeus slightly convex ( Figure 8c ). Antenna with scape 2.24–2.88x and pedicel 1.13–1.57x as long as broad. Flagellum 1.30–1.46x as long as breadth of head. Anellus 0.2–0.5x as long as broad. F1 1.13–1.98x, F2 1.3–1.88x, F3 1.22–1.86x, F4 1.131.74x, F5 1.20–1.71x, F6 1.1–1.6x, F7 1.1–1.42x as long as broad. POL 2.63–4.00x OOL, OOL 0.56–0.89x POD.

Mesosoma ( Figure 8d ). Pronotum 0.83–0.95x as broad as mesoscutum and 0.43–0.63x as long as mesoscutum. Mesoscutellum 1.07–1.2x as long as broad, with frenal area covering about 0.21–0.30x of mesoscutellum length. Metatarsus 0.53–0.68x as long as metatibia. Fore wing 2.60–2.96x as long as broad; marginal vein 3.28–4.33x as long as postmarginal vein and 6.00–8.63x as long as stigmal vein ( Figure 8f ).

Metasoma. Metasoma 0.89–1.2x as long as mesosoma. OI = 5.2–7.6.

Variation (females). Some specimens have entire scapus yellow to light brown, some have scapus yellow laterally; basal part of metafemur of some specimens are yellow to very light yellow. Of the examined 90 females, about 10 specimens vary in number of teeth on hind femora, i.e. some of them are missing the second (the smallest) teeth on right metafemur, some of them are missing the second teeth on left metafemur.

MALE (N = 35): Length of body 2.7–3.26 mm. Similar to females except as follows: head 1.14–1.3x as broad as high; frontovertex 0.78–1.04x as broad as eye height; eye 1.20–1.44x as high as long; parascrobal protuberance 0.16–0.22x as high as head height; malar space 0.48–0.73x as long as breadth of oral fossa and 0.32–0.39x as long as eye height; flagellum longer, 1.43–1.63x as long as breadth of head, pedicel 0.93–1.48x and F7 1.20–1.61x as long as broad; POL 2.24–3.21x OOL, OOL 0.50–0.71x POD; metasoma with yellow subbasal ring extend to 4/5 of metasoma; parascrobal protuberance more raised than in females, about 0.2x as long as eye length (only about 0.13x in females).

Variation (males). There have been reported so called homeomorph and heteromorph (or even their intermediates) males to be found within various genera of Podagrionini (for more details see Delvare 2005). We have found homeomorph and heteromorph males, including some intermediates, within our samples.Most of the examined males have been reared from the same egg case of Archeomantis sp. (lc_11–14, lc_50–52, and lc_130–146, respectively). Homeomorph males are similar to females with slightly different sculpture on propodeum ( Figure 1i ). The intermediates and true heteromorph ( Figure 10a ) males differ in having slightly enlarged all tibiae and mesobasitarus ( Figure 9d ), in number of teeth on metafemur and shape of metatibia ( Figure 9 e–f), in sculpture of propodeum, and shape of petiolus ( Figure 9g, h, j, k ). Within all examined males 5 specimens were true homeomorphs, 20 specimens true heteromorphs, and 10 intermediates.

Distribution. Australia (Queensland, Australian Capital Territory, New South Wales, South Australia, Victoria) and New Caledonia.

Biology. All specimens with known host associations were reared from egg cases of Archimantis sp.

Taxonomic remarks. The comparison of the material preliminarily named morphospecies 1 group A with the type material of the Australian members of the genus Podagrion revealed that morphospecies 1 group A corresponds to P. koebelei Crawford, 1912 and its synonyms P. dolichurum Cockerell, 1930 and P. grotii Girault, 1915 . The remnant of holotype (for holotype condition see below) as well as the non–type material show all generic characters of Lasallegrion . Accordingly, the above redescribed species is transferred into Lasallegrion .

The holotype of L. koebelei lacks head and antennae, the left fore wing is partially covered with glue, the right fore wing and the right hind wing are missing. Unfortunately, the holotype of L. koebelei was damaged during examination for this study, when the plastic card it was mounted on broke at the point where the pin went through. Right hind leg, part of the left hind leg, left fore wing and metasoma were detached from the rest of the body and were mounted on a separate card. The P. grotii type is completely fragmented. It consists of one card–mounted metacoxa, a severely damaged head, antennal parts and a metatibia mounted on a broken slide. It cannot be stated with certainty that all of these fragments actually belong to the same specimen. The type of P. dolichurum is complete and intact. Thus, a complete set of diagnostic characters can only be obtained by combining the types of P. koebelei and P. grotii with the type specimen of P. dolichurum .

Additionally, we identified two new synonyms of L. koebelei , P. holbeini , and P. metatarsum ( Figure 10 b–c).

The three syntypes of P. holbeini (two females and one male mounted on one single card) partially lack heads, but the rest of the body allows us to reliably assign all three specimens to the same species that matches our concept of L. koebelei . The synonymy of P. holbeini with L. koebelei had been suggested already by Bouček (1988).

The five syntypes of P. metatarsum (three females and two males mounted on one single card (SAMA); plus two slides – one (SAMA) with two antennae mounted separately on a slide, labelled ‘ Podagrion metatarsus Gir. , Type ♀ ’; second (QMB) with fragmented head and antennae on it, labelled ‘ Podagrion metatarsus Gir. , Type ♀ ’, 5097) are also partially damaged or fragmented, yet there are enough characters visible that allow assignment to the same species which matches our concept of L. koebelei . Similar to P. holbeini , the synonymy of P. metatarsum with L. koebelei had been suggested already by Bouček (1988).

Although L. koebelei is fairly similar to L. virescens , it can be easily distinguished from the third species of the genus, L. washingtoni , by its very long ovipositor, the funicular segments being always longer than broad, and by the number and shape of the metafemoral teeth. In L. washingtoni the ovipositor is distinctly shorter, the antennal funicle is stouter with its distal segments being quadrate to subquadrate, and the metafemur bears more and smaller ventral teeth than the metafemur of L. koebelei (and L. virescens ). A molecular delimitation of L. koebelei plus L. virescens and L. washingtoni was not possible, because for L. washingtoni no COI sequence data could be obtained.

Lasallegrion koebelei and L. virescens can be differentiated either based on genetic divergence of COI sequences (94.08–94.83%) or morphologically, even if some specimens of both species overlap in some characters, and a combination of all characters is needed to reliably distinguish both species. Furthermore, the multivariate ratio analysis (MRA) resulted in a useful separation of L. koebelei from L. virescens and gave further evidence to separate taxa.

Figures

Figure 1. Maximum likelihood tree. The tree was reconstructed based on COI of Lasallegrion koebelei and L. virescens using RaxML with 1000 bootstrap replications. Only support values ≥50 are shown.

Figure 2. Map of collection sites. Red – Lasallegrion koebelei; green – L. virescens; blue – L. washingtoni (MyMaps by google).

Figure 3. Shape PCA of L. koebelei and L. virescens. The scatterplot shows isosize versus shape PC1.

Figure 4. The scatterplot of best ratios for separating L. koebelei and L. virescens based on the LDA ratio extractor.

Figure 5. Volume renders of the µ-CT scans of Lasallegrion gen. n.; (a), L. virescens, lateral; (b), L. koebelei, lateral. Midsagittal sections showing the inner anatomy. Scale bars: 0.5 mm.

Figure 6. Diagnostic characters of Lasallegrion gen. n.; L. koebelei (a – e), L. washingtoni (f). (a), habitus, lateral; (b), antenna, lateral; (c), head and mesosoma, lateral (arrows indicate pronotal collar carina and metapleuron); (d), propodeum, dorsal; (e), head, posterior; (f), petiolus and first sternite, ventral.

Figure 7. Metepisternum, posteroventral, females. (a), Lasallegrion koebelei; (b), L. virescens; (c), L. washingtoni; (d), Podagrion idomene sp. gr.; (e), P. pachymerum; (f), Mantiphaga gongylusae Risbec.

Figure 8. Lasallegrion koebelei, female. (a), head, dorsal; (b), head, frontal; (c), lower face, detail; (d), mesosoma, dorsal (arrows indicate notauli in their posterior part); (e), right hind leg, outer aspect; (f), left fore wing, ventral.

Figure 9. Lasallegrion koebelei, intermediate (a – c, e, g, h, j), heteromorph (d, f, k) and homeomorph (i) male. (a), head, dorsal; (b), antenna, lateral; (c), head, frontal; (d), left midtibia and midtarsus, outer aspect; (e), right hind leg, outer aspect, intermediate male; (f), right hind leg, outer aspect, heteromorph male; (g), mesoscutellum and propodeum, dorsal; (h), part of propodeum and petiolus, dorsolateral; (i), propodeum and petiolus, dorsal; (j), propodeum and petiolus, dorsal; (k), propodeum and petiolus, dorsal.

Figure 10. (a), Lasallegrion koebelei, heteromorph male habitus, lateral; (b), syntype of P. holbeini, syn. n.; (c), syntype of P. metatarsum, syn. n.; (d), holotype of L. virescens.

Figure 13. (a) – holotype of L. koebelei, lateral aspect; (b) – holotype of L. koebelei, dorsal aspect; (c) – holotype of L. washingtoni, lateral aspect; (d) – holotype of L. washingtoni, dorsal aspect.